Am. J. Pot Res (2010) 87:209217 DOI 10.

1007/s12230-010-9127-6

Potato Flavor
Shelley H. Jansky

Published online: 26 February 2010 # Potato Association of America 2010

Abstract The potato is one of the most popular vegetables worldwide and is the most important vegetable crop in the United States, accounting for nearly one-third of per-capita vegetable consumption. Potatoes can be prepared in many ways, including baking, boiling, roasting, frying, steaming, and microwaving, allowing for a diversity of uses. Most people find potatoes to be an agreeable food and very few actually dislike potatoes. Potato flavor results from the combination of taste, aroma, and texture. Flavor precursors synthesized by the plant are present in raw potatoes and consist mainly of sugars, amino acids, RNA, and lipids. Plant genotype, production environment and storage environment influence the levels of these compounds and the enzymes that react with them to produce flavor compounds. During cooking, flavor precursors react to produce the Maillard reaction compounds and the sugar, lipid and RNA degradation products that contribute to flavor. The identification of major flavor compounds is important for breeders to make gains in selection for enhanced flavor. Resumen La papa es una de las hortalizas ms populares en todo el mundo y es el cultivo hortcola ms importante en los Estados Unidos, ocupando cerca de una tercera parte del consumo per capita de hortalizas. Las papas se pueden preparar de muchas maneras, incluyendo cocinadas, hervidas, rostizadas, fritas, al vapor, y en el horno de microondas, lo cual permite una gran diversidad de usos. La mayora de la gente concuerda en que las papas son un alimento compatible

y a muy poca gente le disgustan las papas. El sabor de la papa resulta de la combinacin de sabor, aroma y textura. Los precursores del sabor sintetizados por la planta estn presentes en las papas crudas y consisten principalmente de azcares, amonocidos, RNA, y lpidos. El genotipo de la planta, el medio ambiente en la produccin y en el almacn influencian los niveles de estos compuestos y a las enzimas que reaccionan con ellos para producir los compuestos del sabor. Durante el cocimiento, los precursores del sabor reaccionan para producir los compuestos de la reaccin de Maillard y para los productos de la degradacin de azcar, lpidos y RNA que contribuyen al sabor. Es importante para los fitomejoradores la identificacin de los principales compuestos del sabor para mejorar en la seleccin de un mayor sabor. Keywords Taste . Texture . Aroma . Potato breeding . Sensory evaluations

Introduction The potato is one of the most popular vegetables worldwide and is the most important vegetable crop in the United States, accounting for nearly one-third of per-capita vegetable consumption (Lin and Yen 2004). Potatoes can be prepared in many ways, including baking, boiling, roasting, frying, steaming, and microwaving, allowing for a diversity of uses. Most people find potatoes to be an agreeable food and less than 1% actually dislike potatoes (Harper 1963). Flavor precursors synthesized by the plant are present in raw potatoes and consist mainly of sugars, amino acids, RNA, and lipids. Plant genotype, production environment and storage environment influence the levels of these compounds and the enzymes that react with them to produce flavor compounds. During cooking, flavor

S. H. Jansky (*) USDA-ARS and Department of Horticulture, University of Wisconsin-Madison, 1575 Linden Drive, Madison, WI 53706, USA e-mail: shelley.jansky@ars.usda.gov

210

Am. J. Pot Res (2010) 87:209217

precursors react to produce the Maillard reaction compounds and the sugar, lipid and RNA degradation products that contribute to flavor (Duckham et al. 2001). The potato industry is interested in the development and promotion of more flavorful potatoes in order to increase consumption. However, potato flavor is not assessed in early generations of breeding programs, where most genetic variability is present. Flavor evaluations based on sensory panels are necessarily low-throughput (Morris et al. 2008). In addition, the optimum flavor goal is typically not known and is likely to be variable among consumer groups. By analyzing flavor only in the advanced stages of a breeding program, breeders are unlikely to make progress toward the development of clones with superior flavor (Wang and Kays 2003). An understanding of the biochemical components of flavor may allow breeders to carry out laboratory analyses in early generations, resulting in the development of advanced selections with enhanced flavor. The three major components of flavor are taste (due to nonvolatile compounds), aroma (due to volatile compounds) and texture (mouthfeel). All three components interact to produce a flavor response. Flavor is strongly influenced not only by potato clone, but also by production and storage environments. This review will address the flavor components of potatoes and the effects of environment on those components.

Taste Human taste receptors monitor bitter, sour, sweet, salty, and umami (a Japanese word meaning delicious). Potato tubers contain all of these taste components except for salty. Unlike fleshy fruits that have evolved to attract dispersal agents, potato tubers have evolved mechanisms to deter consumption by herbivores. Consequently, wild potato tubers may contain bitter-tasting glycoalkaloids to provide protection against pests and disease (Valkonen et al. 1996). Domestication has selected against bitterness in potato tubers, but cultivars still contain some glycoalkaloids (Johns and Alonso 1990). Although the upper limit allowed for a new cultivar is 20 mg/100 g fresh weight, bitterness can be tasted in tubers with glycoalkaloid levels as low as 14 mg/100 g (Sinden et al. 1976). At levels below 10 mg/100 g, glycoalkaloids may make a positive contribution to flavor (Ross et al. 1978). In addition to glycoalkaloids, phenolic compounds contribute to bitterness in potato tubers (Mondy and Gosselin 1988). Since potato skin has higher levels of glycoalkaloids and phenolic compounds, tubers cooked with the skin on are more bitter than those cooked after peeling (Mondy and Gosselin 1988). In addition, when consumed with the skin on, small tubers may taste more bitter than large tubers.

Organic acids in potato tubers can produce a sour or bitter taste. They are produced by the incomplete oxidation of sugars, as well as the deamination of amino acids, ascorbic acid, and polyphenolic acids (Lisinska and Aniolowski 1990). They are generally not considered to be major flavor components (Vainionpaa et al. 2000). However, tubers containing 120 mg/100 g chlorogenic acid were reported to taste slightly sour to some taste panelists (Sinden et al. 1976). It is important to note that taste thresholds for many compounds vary considerably among panel members (Blakeslee and Salmon 1935), so it is not surprising that some panelists detect a sour taste, while others do not. The human genetic variation component of sensory analyses has not been addressed in potato taste panels. Although starch is tasteless, it influences texture and can interact with flavor compounds during cooking (Jitsuyama et al. 2009; Solms and Wyler 1979). Potato tubers also contain low levels of sugars, mainly glucose, fructose, and sucrose, which may contribute directly or indirectly to flavor. Sweetness has historically been considered an undesirable flavor component in potatoes (Burton 1966). However, the consumption of sugars in developed countries has increased in recent decades (Pereira and Simin 2003), so consumers now prefer sweet foods. In fact, we have found that sweetness of baked potatoes is significantly correlated with desirable flavor (Jansky 2008). Sweetness has been reported to be an important component of flavor in other studies as well (Jitsuyama et al. 2009; Lynch and Graven 1971; Vainionpaa et al. 2000; Winfield et al. 2005). Potato chips with added sugar were also preferred by consumers (Maier et al. 2007). Ribonucleotides act as precursors for umami compounds, which act as flavor enhancers. Although low in raw potatoes, relatively high levels of 5 ribonucleotides are released by the enzymatic hydrolysis of RNA as tubers are heated during cooking. Nucleases are especially active at around 50C, so high activity is expected during early stages of cooking (Solms and Wyler 1979). Steamed or boiled tubers of Phureja Group cultivars with higher levels of glutamate and guanosine 5-monophosphate (GMP) than Tuberosum Group cultivars had higher acceptability scores in taste tests (Morris et al. 2007, 2008). The most important ribonucleotides for flavor enhancement are inosine 5monophosphate and GMP. Both levels and types of ribonucleotides vary among potato cultivars, probably due to differences in ribonucleases (Maga and McNeill 1986). A ribonuclease with an enhanced level of expression was found in flavorful Phureja Group clones (Ducreux et al. 2008). A synergistic effect is detected when 5 ribonucleotides interact with amino acids, especially glutamate. In fact, the products of interactions between amino acids and 5 ribonucleotides are considered to be mainly responsible

Am. J. Pot Res (2010) 87:209217

211

for boiled potato flavor (Halpern 2000; Solms 1971). Sugars may also contribute to umami taste characters in the form of glutamate glycoconjugates (Beksan et al. 2003). In addition, potassium salts have been found to enhance umami taste intensity (Ugawa and Kurihara 1994). However, significant levels of potassium leach out of potatoes during boiling, which may reduce opportunities for this interaction (Bethke and Jansky 2008). Cooking oil is an important taste component of processed potato products. Frying oils derived from different sources have been reported to impart unique flavors during processing (Weiss 1983). However, in another study, sensory evaluations could not detect a major effect of cooking oil on chip flavor (Roberson et al. 1978). In addition, while potato chip flavor intensity was not found to depend on oil type, intensities of rancidity and off-flavor after storage did vary (Pangloli et al. 2002). Color is not directly a taste component of flavor, but it influences the perception of taste (Moriera et al. 1999). For example, deep orange sweet potatoes were described by taste panelists as pumpkin- or carrot-like (McLaurin and Jays 1992). Sensory analyses of colored flesh potatoes have not yet been published. Color has always been important in fried products, though, since Maillard reaction products formed at high processing temperatures result in unacceptably dark chips (Denny and Thornton 1940; Habib and Brown 1957). In the Maillard reaction, reducing sugars (glucose and fructose) interact with amino acids at high temperature to produce dark-colored products. Industry standards require light-colored chips, but blindfolded taste panelists preferred the taste and odor of dark-colored chips,

presumably due to Maillard reaction products that make a contribution as flavor compounds (Maga 1973).

Aroma Unlike fleshy fruits, potato tubers do not emit volatile flavor compounds at maturity. Instead, aroma compounds develop when tuber tissues are sliced and/or heated. Cooked potatoes produce hundreds of aroma compounds (Coleman et al. 1981), which exhibit a wide range of concentrations and odor thresholds. Several methods are used for the analysis of aroma profiles of potato products, including simultaneous distillation and extraction (SDE), solvent-assisted flavor evaporation (SAFE), and solid-phase microextraction (SPME) (Majcher and Jelen 2009). The most important aroma compounds are produced by lipid degradation and by the Maillard reaction and/or sugar degradation during heating (Oruna-Concha et al. 2002). In potato chips, the most abundant aroma compounds are the oxidation products of the fatty acids in the frying oil (Pangloli et al. 2002). Sugars and amino acids are also important for the development of flavor compounds in chips (Martin and Ames 2001). A summary of important cooked potato aroma compounds is presented in Table 1. During baking, the tuber warms first at the surface and water evaporates from the skin. Over time, a crust develops and the tuber gradually warms from the outside toward the interior. At high baking temperatures, tubers produce a complex array of volatile compounds, including lipid degradation products, Maillard reaction products, sulfur

Table 1 Compounds reported to have a strong impact on cooked potato flavor Compound 2,-4-decadienal c4-heptenal 1-octen-3-ol alkyl pyrazines Origin lipid degradation lipid degradation lipid degradation Maillard reaction/sugar degradation Maillard reaction/sugar degradation Maillard reaction/sugar degradation Maillard reaction/sugar degradation raw tuber raw tuber/soil microbes sulfur amino acid sulfur amino acid Flavor earthy, fatty cold boiled potato earthy, mushroom earthy, roasted, nutty, buttery fruity, fatty, floral fruity green fruity, floral musty, earthy cooked potato cooked onion References Oruna-Concha et al. 2001 Josephson and Lindsay 1987 Oruna-Concha et al. 2001 Maga and Holm 1992; Oruna-Concha et al. 2001; Coleman and Ho 1980; Buttery et al. 1971 Duckham et al. 2001, 2002 Duckham et al. 2001, 2002 Oruna-Concha et al. 2001 Duckham et al. 2001 Duckham et al. 2001, 2002 Duckham et al. 2002; Oruna-Concha et al. 2001 Duckham et al. 2001, 2002; Oruna-Concha et al. 2001

decanal 2- and 3-methylbutanal phenylacetaldehyde terpenes (-copaene, damascenone, others) methoxypyrazines methional dimethyl trisulfide, dimethyl disulfide

212

Am. J. Pot Res (2010) 87:209217

compounds, and methoxypyrazines (Oruna-Concha et al. 2001). Pyrazines produced by the Maillard reaction are considered to be among the most important and characteristic components of baked potato flavor (Buttery et al. 1973). There is a strong positive relationship between pyrazines and organoleptic quality in both baked potatoes (Maga and Holm 1992) and potato chips (Maga and Sizer 1973). In contrast to baked potatoes, water loss in boiled potatoes is minimal and the interior warms quickly but the tuber temperature never exceeds 100C. The major aroma components of boiled potatoes are derived from lipid degradation, the Maillard reaction, and the Strecker degradation of methionine (Maga 1994; Mutti and Grosch 1999; Taylor et al. 2007; Ulrich et al. 2000). Boiled tubers contain higher levels of lipid degradation products than do baked potatoes (Oruna-Concha et al. 2002). The disruption of tissues during slicing in preparation for boiling provides more opportunities for lipoxygenase to come into contact with its substrates. Fatty acids degrade to produce aldehydes and ketones, which contribute to fatty, fruity, and floral flavor notes (Duckham et al. 2002). Another major contributor to boiled potato aroma is the lipid oxidation product c4-heptanal, which produces an earthy aroma at low levels (Josephson and Lindsay 1987). However, samples tasted stale when high levels of c4-heptanal were added to mashed potatoes. Similarly, lipid oxidation products have also been implicated in the off-flavor of potato flakes (Gosset et al. 2008) and boiled potatoes after they are refrigerated (Peterson and Poll 1999). These studies illustrate the concept that high levels of flavor components are not always desirable. It is necessary to determine the upper and lower limits of these components with respect to acceptable flavor contributions. During microwave baking, tuber temperature increases relatively uniformly, with all parts reaching 100C within a few minutes of each other (Oruna-Concha et al. 2002). In contrast to oven baking, the skin remains cooler than the interior of the tuber, due to evaporative cooling. A crust does not develop, so the rate of water loss is higher. Microwave-baked potatoes have lower levels of volatiles than oven-baked or boiled potatoes, probably due to evaporative cooling at the tuber surface and the loss of volatile compounds through co-distillation as water evaporates. Consequently, they tend to be more bland than boiled or baked potatoes, and they receive lower ratings in sensory analyses (Brittin and Trevino 1980; Maga and Twomey 1977). Methional is a major aroma compound that is formed by the Strecker degradation reaction, in which intermediates in the Maillard reaction interact with the amino acid methionine. Levels of methional vary among cultivars and production environments (Duckham et al. 2002; OrunaConcha et al. 2001). This compound is not detectable in all

potato cultivars. In one study, it was found in only five out of 11 cultivars evaluated (Duckham et al. 2001). Unlike most flavor compounds, methoxypyrazines are present in raw tubers. They may be produced by the tuber or by soil bacteria (Pseudomonas taetrolens) and then absorbed by the tuber (Buttery et al. 1973). Methyoxypyrazine levels vary with food preparation methods. They have been found in higher amounts in boiled potatoes than in baked potatoes, possibly due to cell damage that results from peeling tubers in preparation for boiling (Oruna-Concha et al. 2002). The odor threshold of methoxypyrazines is very low. For example, the odor threshold of a common methoxypyrazine, 2-methoxy-3-isopropylpyrazine, in water is 2 parts in 1012. Consequently, methoxypyrazines are said to have a high aroma impact value (Duckham et al. 2002). 2-methoxy-3isopropylpyrazine has been found in tubers at 1 part in 1010 (Buttery et al. 1973). In biochemical analyses, methoxypyrazines are not detectable in many cultivars and are present at very low levels in others. Because of their high aroma impact values, though, small changes in methoxypyrazine levels are expected to have large effects on flavor. Pigments such as anthocyanins and carotenoids are not thought to directly influence flavor. However, carotenoid degradation products following cooking have been reported to contribute to flavor of sweetpotato (Wang and Kays 2000). These compounds have low odor thresholds, so they have the ability to make significant contributions to aroma. Breeders are interested in creating high antioxidant clones for their increased nutritional value, but this may have a negative effect on flavor. Increasing the terpene biosynthetic pathway that leads to increased levels of carotenoids in sweetpotatoes resulted in new flavors that consumers generally found disagreeable (Wang and Kays 2003). As a result of an expression profiling study comparing flavorful Phureja Group clones with bland Tuberosum Group clones, the sesquiterpene -copaene was found to contribute positively to flavor (Ducreux et al. 2008). The gene encoding -copaene synthase is expressed at a higher level in Phureja Group clones. It is interesting to note that a number of sesquiterpenes are produced by tubers in response to infection by pathogens (Engstrom et al. 1999; Hildenbrand and Ninnemann 1994).

Texture Texture is one of the most important quality attributes of potato tubers. It not only affects consumer preference, but it also influences the release of volatile flavor components during chewing (Lucas et al. 2002). This component of flavor is easily recognizable by consumers, who tend to have distinct preferences. Potato texture is a complex trait, controlled by many factors, including dry matter content,

Am. J. Pot Res (2010) 87:209217

213

specific gravity, amylose, sugars, protein, and nitrogen levels in tubers (Arvanitoyannis et al. 2008; Jitsuyama et al. 2009). However, much variation can be explained by determining the degree of a tubers mealiness or, at the opposite end of the spectrum, waxiness (Faulks and Griffiths 1983; van Marle et al. 1997b). A mealy potato is dry and granular, while a waxy potato is moist and gummy. Mealiness has been found to be associated with high dry matter content (Jansky 2008; Leung et al. 1983; Lynch and Graven 1971; van Dijk et al. 2002). Culinary uses of potato cultivars relate to their texture, with those developed for boiling retaining their shape during cooking (Smith et al. 2009). Mealiness is one of the readily described components of flavor for taste panelists. In fact, detection thresholds for mealiness by taste panelists have been identified (Murphy et al. 1967). However, dry matter content does not always explain mealiness (Brittin and Trevino 1980). In one sensory analysis study, the cultivar Ontario was judged to be less mealy than other cultivars in the trial, even though its dry matter content was similar (True and Work 1981). Another component of texture is the size and structure of starch grains in raw tuber tissue (Thybo and Martens 1999). Starch gelatinizes during cooking and creates pressure in cells as it expands. The proportion of each tuber cell occupied by gelatinized starch influences tuber moistness (Martens and Thybo 2000). Cells that are filled with gelatinized starch are associated with a mealy texture, while those with less starch and more loosely-held water produce a waxy texture (Martens and Thybo 2000; McComber et al. 1994). The loosely held water in the latter cell type is released upon chewing, producing a moist mouthfeel. The gelatinized starch in the mealy types retains water, creating a dry mouthfeel. Cell size has also been found to be associated with mealiness (Barrios et al. 1963). Tubers given high mealiness scores by taste panelists were found to contain more starch and have larger cells than less mealy tubers. Ploidy effects on cell size may influence texture. Diploid clones were found to have a more floury texture than related tetraploid clones (De Maine et al. 1993). Cell wall characteristics can also affect texture (Jarvis and Duncan 1992; Jarvis et al. 1992; Martens and Thybo 2000; McComber et al. 1994; van Marle et al. 1997b). Pectin methyl esterase activity is important for creating firm tissue during cooking by cross-linking pectin in the middle lamella (Faulks and Griffiths 1983; Thybo and Martens 1999). Cultivars vary in cell wall density and the degree of solubilization of the middle lamella and cell walls, which influence texture (van Marle et al. 1997b). The mealy cultivar Irene was found to have more cell wall material per unit cell surface area than the nonmealy cultivar Nicola. The cell wall and middle lamella were also found to be thicker in the mealy cultivar Russet Burbank than in the

waxy cultivar Red Pontiac (McComber et al. 1994). The thicker cell walls and middle lamellae, and stronger pectic substances may be more resistant to shearing, resulting in a hard, particulate mouthfeel. Similarly, mealiness was reported to correlate with cell cohesiveness and adhesiveness (Leung et al. 1983). In contrast to the soft texture of baked or boiled potatoes, fried potatoes must be crispy. French fries should have a crispy exterior, while chips should be crispy throughout. Crust formation results from the softening of the middle lamella and the gelatinization and dehydration of starch during heating (Troncoso et al. 2009). While high dry matter is important to reduce the oil content of chips, if the dry matter is too high (>25%), then the chips will have a hard texture (Kita 2002). Potato chip texture depends mainly on starch composition, but also on non-starch polysaccharides, lignin, and protein nitrogen (Kita 2002).

Effect of Production Environment on Flavor The effect of production environment on sensory quality can be large and may sometimes be more important than that of cultivar (Arvanitoyannis et al. 2008; Faulks and Griffiths 1983; Jansky 2008). Levels of methional vary among tubers harvested from different production environments (Duckham et al. 2002; Oruna-Concha et al. 2001). Because methionine contains sulfur, sulfur application rates in the field may account for some differences in methional levels (Duckham et al. 2002). Similarly, potassium application in the field may influence umami flavor intensity (Morris et al. 2007). Because methoxypyrazines may be produced by soil bacteria, soil microbe populations may influence flavor (Buttery et al. 1973). Levels of off-flavors may be influenced by production environment as well. As nitrogen levels increase, sensory quality decreases, probably due to the production of acrid tasting amides and amines (Cieslik 1997; Jansky 2008; Thybo et al. 2006). Differences in production environment were found to influence the development of off-flavor in precooked vacuum-packed potatoes (Jensen et al. 1999). In another study, a musty off-flavor detected in cooked potatoes was found to be due to the presence of 2,4,6-trichloroanisole. While this compound is not a known metabolite of potato, it was found in tubers harvested from pesticide-treated soils after an exceptionally warm production year (Daniels-Lake et al. 2007). Glycoalkaloid levels and, consequently, bitter flavor, may increase in tubers grown under stressful conditions and in tubers that are exposed to light (Percival et al. 1994; Sinden and Webb, 1972; Uppal 1987; Valkonen et al. 1996). In addition, bruising during harvest can result in significant increases in levels of glycoalkaloids and chlorogenic acid (Dale et al. 1998).

214

Am. J. Pot Res (2010) 87:209217

Some studies have evaluated the effects of organic versus conventional production systems on sensory attributes. Taste panelists were able to distinguish between conventionally and organically grown boiled red potatoes if the skin was left on the tubers while boiling (Wszelaki et al. 2005). However, if the skin was removed during boiling, then differences could not be detected. Moistness, mealiness, color, and odor were reported to be influenced to a small extent by organic fertilizer treatments (Thybo et al. 2002). However, organic production systems were not found to be consistently superior or inferior to conventional (Hajslova et al. 2005; Jansky 2008; Woese et al. 1997).

Effect of Storage Environment on Flavor Changes in sensory quality occur during the storage of potato tubers. In a sensory analysis study, stored tubers were found to be mealier, sweeter, and more flavorful than fresh tubers (Jansky 2008). Tuber pH, dry matter content, and levels of off-flavor have also been reported to decrease during storage (Arvanitoyannis et al. 2008; Jansky 2008; Thybo et al. 2006). Russet Burbank ranked high and Ontario ranked low for flavor preference in fresh baked potatoes, but differences were not detected in stored tubers (True and Work 1981). In contrast to fresh potatoes, processed potatoes may decrease in quality when stored. Potato chips exhibit a deterioration in flavor over time (Roberson et al. 1978). Chips stored in the light deteriorate more quickly than those in the dark (Dornseifer and Powers 1965). Because sprout inhibitors are commonly used to maintain tuber quality during storage, their potential effects on flavor are important. Taste panelists were not able to detect the sprout inhibitor CIPC (isopropyl-Nchlorophenyl carbamate), but they were able to taste residual levels of the alternative sprout inhibitor 1,8cineole (Boylston et al. 2001). When potato tubers are cooked, fatty acids degrade to produce aldehydes and ketones, which contribute to flavor (Duckham et al. 2002). Total levels of fatty acids increase during storage and contribute to the aroma component of both baked potatoes (Duckham et al. 2002) and potato chips (Dornseifer and Powers 1965). Tubers alter their fatty acid profiles as they acclimate to cold storage temperatures. Consequently, both levels and types of fatty acids change. Cultivars vary in the way they change their fatty acid profiles during cold storage (Mondy et al. 1963). However, levels of linoleic acid typically decrease and -linolenic acid increases (Dobson et al. 2004). Lipoxygenase activity in potato tubers increases during cold storage. The concentrations of off-flavor products, such as aldehydes, were found to decrease when these tubers were boiled and then chilled (Peterson et al. 2003).

The types and levels of Maillard reaction components of flavor also change during storage, due to alterations in enzyme activities and levels of flavor precursors, such as sugars (Duckham et al. 2002). Levels of reducing sugars and free amino acids increase during cold storage (Blenkinsop et al. 2002; Fitzpatrick and Porter 1966; Sowokinos 2001). Consequently, aroma intensity due to pyrazines increases during cold storage (Duckham et al. 2002). In addition, Maillard-derived glutamate glycoconjugates, which act as umami compounds, are expected to increase with the accumulation of sugars during storage. Cultivar-dependent effects of storage on texture have been reported (Faulks and Griffiths 1983; Martens and Thybo 2000; Thybo and Martens 1999). In general, though, tuber mealiness decreases during storage (Ridley and Lindsay 1984; Shetty et al. 1992; van Marle et al. 1997a). Starch is broken down during storage, leading to pitted starch granules (Cottrell et al. 1993). The degradation of starch and the breakdown of the middle lamella during storage may lead to less mealy tubers (Martens and Thybo 2000). In addition, potato tubers lose water during storage and become less turgid. As a result, cells lose their elasticity and are ruptured during cooking, resulting in a less mealy texture (Shetty et al. 1992).

Conclusions Flavor is an important marketing trait for any food crop, but potato breeders do not focus on selection for superior flavor. The development of potato varieties with enhanced flavor has the potential to increase consumer interest in the consumption of potatoes. Breeding progress for improved flavor requires both genetic diversity for the trait and an effective way to identify superior clones. The first requirement may be the lesser of the two potential constraints because tremendous diversity is found in cultivated and wild relatives of potato. The best source of valuable flavor diversity is likely to be in Andean land races, which were selected in regions of high genetic diversity. Flavor was probably an important selection criterion in this germplasm, because the indigenous people who developed them consumed large quantities of potatoes and had access to tremendous genetic diversity. In fact, Phureja Group potatoes have been found to be good sources of flavorful potatoes (De Maine et al. 1993; Winfield et al. 2005). After the major biochemical components of flavor are identified, it would be valuable to screen wild Solanum species for exceptional levels of those components. The second requirement is more problematic because flavor evaluation based on sensory panels is extremely time consuming and can be carried out on only a few samples at a time. Breeders must to be able to characterize specific components of

Am. J. Pot Res (2010) 87:209217

215 potatoes treated with CIPC and alternative sprout inhibitors. American Journal of Potato Research 78: 99107. Brittin, H.C., and J.E. Trevino. 1980. Acceptability of microwave and conventionally baked potatoes. Journal of Food Science 45: 14251427. Burton, W.G. 1966. The potato H. Wageningen: Veenan and Zonen. Buttery, R.G., R.M. Seifert, D.G. Guadagni, and L.C. Ling. 1971. Characterization of volatile pyrazine and pyridine components of potato chips. Journal of Agricultural and Food Chemistry 19: 969971. Buttery, R.G., D.G. Guadagni, and L.C. Ling. 1973. Volatile components of baked potatoes. Journal of the Science of Food and Agriculture 24: 11251131. Cieslik, E. 1997. Effect of the levels of nitrate and nitrites on the nutritional and sensory quality of potato tubers. Hygiene and Nutrition in Foodservice and Catering 1: 225230. Coleman, E.C., and C.-T. Ho. 1980. Chemistry of baked potato flavor. 1. Pyrazines and Thiazoles identified in the volatile flavor of baked potato. Journal of Agricultural and Food Chemistry 28: 6870. Coleman, E.C., C.-T. Ho, and S.S. Chang. 1981. Isolation and identification of volatile compounds from baked potatoes. Journal of Agricultural and Food Chemistry 29: 4248. Cottrell, J.E., C.M. Duffus, G.R. MacKay, and M.J. Allison. 1993. Changes in the surface morphology of starch granules of the cultivated potato, Solanum tuberosum L. during storage. Potato Research 36: 119125. Dale, M.F.B., D.W. Griffiths, and H. Bain. 1998. Effect of bruising on the total glycoalkaloid and chlorogenic acid content of potato (Solanum tuberosum) tubers of five cultivars. Journal of the Science of Food and Agriculture 77: 499505. Daniels-Lake, B.J., R.K. Prange, S.O. Gaul, K.B. McRae, R. de Antueno, and D. McLachlan. 2007. A musty off flavor in Nova Scotia potatoes is associated with 2, 4, 6-trichloroanisole released from pesticide-treated soils and high soil temperature. Journal of the American Society for Horticultural Science 132: 112119. De Maine, M., C.P. Carroll, and C.J.W. Torrance. 1993. Culinary quality of tubers derived from Solanum phureja and S. tuberosum x S. phureja hybrids. Journal of Agricultural Science 120: 213 217. Denny, F.E., and N.C. Thornton. 1940. Factors for color in the production of potato chips. Contributions from the Boyce Thompson Institute 11: 291302. Dobson, G., D.W. Griffiths, H.V. Davies, and J.W. McNicol. 2004. Comparison of fatty acid and polar lipid contents of tubers from two potato species, Solanum tuberosum and Solanum phureja. Journal of Agricultural and Food Chemistry 52: 63066314. Dornseifer, T.P., and J.J. Powers. 1965. Volatile constituents of potato chips and changes during storage. Food Technology 19: 877879. Duckham, S.C., A.T. Dodson, J. Bakker, and J.M. Ames. 2001. Volatile flavour components of baked potato flesh: A comparison of eleven potato cultivars. Nahrung/Food 45: 317323. Duckham, S.C., A.T. Dodson, J. Bakker, and J.M. Ames. 2002. Effect of cultivar and storage time on the volatile flavor components of baked potato. Journal of Agricultural and Food Chemistry 50: 56405648. Ducreux, L.J.M., W.L. Morris, I.M. Prosser, J.A. Morris, M.H. Beale, F. Wright, T. Shepherd, G.J. Bryan, P.E. Hedley, and M.A. Taylor. 2008. Expression profiling of potato germplasm differentiated in quality traits leads to the identification of candidate flavour and texture genes. Journal of Experimental Botany 59: 42194231. Engstrom, K., A.K. Widmark, S. Brishammar, and S. Helmersson. 1999. Antifungal activity to Phytophthora infestans of sesquiterpenoids from infected potato tubers. Potato Research 42: 4350. Faulks, R.M., and N.M. Griffiths. 1983. Influence of variety, site and storage on physical, sensory and compositional aspects of

flavor to allow them to identify superior clones in their programs. In addition, the effects of production environment and storage environment on flavor must be understood. While the biochemical components of potato flavor are well-characterized, the connection to taste panel data is often missing. In addition, taste and aroma compounds vary in their relative impact on overall flavor. Therefore, concentrations of biochemical components may not predict their effect on flavor (Wang and Kays 2003). Despite these complications, efforts have been made to associate large scale biochemical analyses with flavor. One strategy is to find a few major contributors to flavor by comparing phenotypically different clones and then focus on those compounds during selection (Beckmann et al. 2007; Ducreux et al. 2008). Another strategy is to develop flavor profiles of standard cultivars using a large biochemical data set and then characterize clones in segregating populations based on their relative positions to the standard cultivars (Wang and Kays 2003). This would identify clones that are similar to conventional cultivars, as well as clones with novel flavor profiles. It might also be possible to identify flavors preferred by different types of consumers, such as ethnic or age groups, and select clones based on target classes.

References
Arvanitoyannis, I.S., O. Vaitsi, and A. Mavromatis. 2008. Physicochemical and sensory attributes in conjunction with multivariate analysis of two potato (Solanum tuberosum L.) cultivars after 90 days of storage: An exploratory authentication study. International Journal of Food Science and Technology 43: 19601970. Barrios, E.P., D.W. Newsom, and J.C. Miller. 1963. Some factors influencing the culinary quality of Irish potatoes. II. Physical characters. American Potato Journal 40: 200208. Beckmann, M., D.P. Enot, D.P. Overy, and J. Draper. 2007. Representation, comparison, and interpretation of metabolome fingerprint data for total composition analysis and quality trait investigation in potato cultivars. Journal of Agricultural and Food Chemistry 55: 34443451. Beksan, E., P. Schieberle, F. Robert, I. Blank, L.B. Fay, H. Schlichtherle-Cerny, and T. Hofmann. 2003. Synthesis and sensory characterization of novel umami-tasting glutamate glycoconjugates. Journal of Agricultural and Food Chemistry 51: 54285436. Bethke, P.C., and S. Jansky. 2008. The effects of boiling and leaching on the content of potassium and other minerals in potatoes. Journal of Food Science 73: H80H85. Blakeslee, A.F., and T.N. Salmon. 1935. Genetics of sensory thresholds: Individual taste reactions for different substances. Proceedings of the National Academy of Sciences 21: 8490. Blenkinsop, R.W., L.J. Copp, R.Y. Yada, and A.G. Marangoni. 2002. Changes in compositional parameters of tubers of potato (Solanum tuberosum) during low-temperature storage and their relationship to chip processing quality. Journal of Agricultural and Food Chemistry 50: 45454553. Boylston, T.D., J.R. Powers, K.M. Weller, and J. Yang. 2001. Comparison of sensory differences of stored Russet Burbank

216 mashed potato. Journal of the Science of Food and Agriculture 34: 979986. Fitzpatrick, T.J., and W.L. Porter. 1966. Changes in the sugars and amino acids in chips made from fresh, stored and reconditioned potatoes. American Potato Journal 43: 238248. Gosset, V., C. Gobel, G.L. Laine, P. Delaplace, P. du Jardin, I. Feussner, and M.-L. Fauconnier. 2008. The role of oxylipins and antioxidants on off-flavor precursor formation during potato flake processing. Journal of Agricultural and Food Chemistry 56: 1128511292. Habib, A.T., and H.D. Brown. 1957. Role of reducing sugars and amino acids in the browning of potato chips. Food Technology 11: 5859. Hajslova, J., V. Schulzova, P. Slanina, K. Janne, K.E. Hellenas, and C. Andersson. 2005. Quality of organically and conventionally grown potatoes: Four-year study of micronutrients, metals, secondary metabolites, enzymic browning, and organoleptic qualities. Food Additives and Contaminants 22: 514534. Halpern, B.P. 2000. Glutamate and the flavor of foods. Journal of Nutrition 130: 910s914s. Harper, R. 1963. Some attitudes to vegetables, and their implications. Nature 200: 1418. Hildenbrand, S., and H. Ninnemann. 1994. Kinetics of phytoalexin acumulation in potato tubers of different genotypes infected with Ervinia carotovora ssp. atroseptica. Physiological and Molecular Plant Pathology 44: 335347. Jansky, S. 2008. Genotypic and environmental contributions to baked potato flavor. American Journal of Potato Research 85: 455465. Jarvis, M.C., and H.J. Duncan. 1992. The textural analysis of cooked potatoes. 1. Physical properties of the separate measurement of softness and dryness. Potato Research 35: 8391. Jarvis, M.C., E. MacKenzie, and H.J. Duncan. 1992. The textural analysis of cooked potatoes. 2. Swelling pressure of starch during gelatinisation. Potato Research 35: 93102. Jensen, K., M.A. Petersen, L. Poll, and P.B. Brockhoff. 1999. Influence of variety and growing location on the development of off-flavor in precooked vacuum-packed potatoes. Journal of Agricultural and Food Chemistry 47: 11451149. Jitsuyama, Y., A. Tago, C. Mizukami, K. Iwama, and S. Ichkawa. 2009. Endogenous components and tissue cell morphological traits of fresh potato tubers affect the flavor of steamed tubers. American Journal of Potato Research 86: 430441. Johns, T., and J.G. Alonso. 1990. Glycoalkaloid change during the domestication of the potato, Solanum Section Petota. Euphytica 50: 203210. Josephson, D.B., and R.C. Lindsay. 1987. C4-heptenal: an influential volatile compound in boiled potato flavor. Journal of Food Science 52: 328331. Kita, A. 2002. The influence of potato chemical composition on crisp texture. Food Chemistry 76: 173179. Leung, H.K., F.H. Barron, and D.C. Davis. 1983. Textural and rheological properties of cooked potatoes. Journal of Food Science 48: 14701474. Lin, B.-H., and S.T. Yen. 2004. U.S. potato consumption: Looking ahead to 2020. Journal of Food Products Marketing 10: 4965. Lisinska, G., and K. Aniolowski. 1990. Organic acids in potato tubers: Part 1The effect of storage temperatures and time on citric and malic acid contents of potato tubers. Food Chemistry 38: 255261. Lucas, P.W., J.F. Prinz, K.R. Agrawal, and I.C. Bruce. 2002. Food physics and oral physiology. Food Quality and Preference 13: 203213. Lynch, D.R., and E.H. Graven. 1971. Effect of rate of fertilization and stage of harvest on yield and culinary quality of four potato cultivars. Agroplantae 3: 714. Maga, J.A. 1973. Influence of freshness and color on potato chip sensory preferences. Journal of Food Science 38: 12511252.

Am. J. Pot Res (2010) 87:209217 Maga, J.A. 1994. Potato Flavor. Food Reviews International 10: 148. Maga, J.A., and D.G. Holm. 1992. Subjective and objective comparison of baked potato aroma as influenced by variety/clone. In Food science and human nutrition, ed. G. Charalambous, 537 541. Amsterdam: Elsevier Science Publishers. Maga, J.A., and J.G. McNeill. 1986. Nucleotide composition of incubated potatoes from five potato (Solanum tuberosum) cultivars. Lebensmittel-Wissenschaft Technologie 19: 3133. Maga, J.A., and C.E. Sizer. 1973. Pyrazines in foods. A review. Journal of Agricultural and Food Chemistry 21: 2230. Maga, J.A., and J.A. Twomey. 1977. Sensory comparison of four potato varieties baked conventionally and by microwaves. Journal of Food Science 42: 541542. Maier, A., Z. Vickers, and J.J. Inman. 2007. Sensory-specific satiety, its crossovers, and subsequent choice of potato chip flavors. Appetite 49: 419. Majcher, M., and H.H. Jelen. 2009. Comparison of suitability of SPME, SAFE and SDE methods for isolation of flavor compounds from extruded potato snacks. Journal of Food Composition and Analysis 22: 606612. Martens, H.J., and A.K. Thybo. 2000. An integrated microstructural, sensory and instrumental approach to describe potato texture. Lebensmittel-Wissenschaft and Technologie 33: 471482. Martin, F.L., and J.M. Ames. 2001. Formation of Strecker aldehydes and pyrazines in a fried potato model system. Journal of Agricultural and Food Chemistry 49: 38853892. McComber, D.R., H.T. Horner, M.A. Chamberline, and D.F. Cox. 1994. Potato cultivar differences associated with mealiness. Journal of Agricultural and Food Chemistry 42: 24332439. McLaurin, W.K., and S.J. Jays. 1992. Genetic diversity in sweetpotato flavor. In The Sweetpotato in the 21st Century. 2nd International Symposium on Sweetpotatoes, eds. W.A. Hill, et al., 420427. Tuskegee, AL. Mondy, N.I., and B. Gosselin. 1988. Effect of peeling on total phenols, total glycoalkaloids, discoloration and flavor of cooked potatoes. Journal of Food Science 53: 756759. Mondy, N., L.R. Mattick, and E. Owens. 1963. The effects of storage on total lipides and the fatty acid composition of potatoes. Agricultural and Food Chemistry 11: 328329. Moriera, G., M.E. Castell-Perez, and M.A. Barrufet. 1999. Deep fat frying: Fundamentals and applications. New York: Aspen Publishers, Inc. Morris, W.L., H.A. Ross, J.M. Ducreux, J. Bradshaw, G.J. Bryan, and M.A. Taylor. 2007. Umami compounds are a determinant of the flavor of potato (Solanum tuberosum L.). Journal of Agricultural and Food Chemistry 55: 96279633. Morris, W.L., L.J.M. Ducreux, G.J. Bryan, and M.A. Taylor. 2008. Molecular dissection of sensory traits in the potato tuber. American Journal of Potato Research 85: 286297. Murphy, E.F., R.H. True, and J.M. Hogan. 1967. Detection threshold of sensory panels for mealiness of baked potatoes as related to specific gravity differences. American Potato Journal 44: 442451. Mutti, B., and W. Grosch. 1999. Potent odorants of boiled potatoes. Nahrung/Food 43: 302306. Oruna-Concha, M.J., C. Duckham, and J.M. Ames. 2001. Comparison of volatile compounds isolated from the skin and flesh of four potato cultivars after baking. Journal of Agricultural and Food Chemistry 49: 24142421. Oruna-Concha, M.J., J. Bakker, and J.M. Ames. 2002. Comparison of the volatile components of two cultivars of potato cooked by boiling, conventional baking and microwave baking. Journal of the Science of Food and Agriculture 82: 10801087. Pangloli, P., S.L. Melton, J.L. Collins, M.P. Penfield, and A.M. Saxton. 2002. Flavor and storage stability of potato chips fried in cottonseed and sunflower oild and palm olein/sunflower blends. Food Chemistry and Toxicology 67: 97103.

Am. J. Pot Res (2010) 87:209217 Percival, G.C., G.R. Dixon, and A. Sword. 1994. Glycoalkaloid concentration of potato tubers following continuous illumination. Journal of the Science of Food and Agriculture 66: 139144. Pereira, M.A., and L. Simin. 2003. Types of carbohydrates and risk of cardiovascular disease. Journal of Womens Health 12: 115122. Peterson, M.A., and L. Poll. 1999. Identification of compounds contributing to boiled potato off-flavour (POF). LebensmittleWissenschaft Technologie 32: 3240. Peterson, M.A., L. Poll, and L.M. Larsen. 2003. Changes in flavoraffecting aroma compounds during potato storage are not associated with lipoxygenase activity. American Journal of Potato Research 80: 397402. Ridley, S.C., and R.C. Lindsay. 1984. Evaluation of factors affecting predictability of mealy textures in potatoes. Journal of Food Quality 7: 116. Roberson, J.A., W.H. Morrison, B.G. Lyon, and R.L. Shaw. 1978. Flavor and chemical evaluation of potato chips fried in sunflower, cottonseed, and palm oils. Journal of Food Science 43: 420423. Ross, H., P. Pasemann, and W. Nitzsche. 1978. Glycoalkaloid content of potatoes and its relationship to location, year and taste. Zeitschrift fr Pflanzenzchtung 80: 6479. Shetty, K.K., R.B. Dwelle, and J.K. Fellman. 1992. Sensory and cooking quality of individually film wrapped potatoes. American Potato Journal 69: 275287. Sinden, S.L., and R.L. Webb. 1972. Effect of variety and location on the glycoalkaloid content of potatoes. American Potato Journal 49: 334338. Sinden, S.L., K.L. Deahl, and B.B. Aulenbach. 1976. Effect of glycoalkaloids and phenolics on potato flavor. Journal of Food Science 41: 520523. Smith, M.F., C.S. Leighton, L. Morey, and H.C. Schonfeldt. 2009. Application of partial least squares regression to relate tastiness of boiled potatoes to chemical and physical tests. South African Journal of Plant and Soil 26: 3135. Solms, J. 1971. Nonvolatile compounds and flavour. London: Academic. Solms, J., and R. Wyler. 1979. Taste components of potatoes. In Food taste chemistry, ed. J.C. Boudreau, 175184. Washington: American Chemical Society. Sowokinos, J.R. 2001. Biochemical and molecular control of coldinduced sweetening in potatoes. American Journal of Potato Research 78: 221236. Taylor, M.A., G.J. McDougall, and D. Stewart. 2007. Potato flavor and texture. In Potato biology and biotechnology, ed. Vreugdenhil, 525540. San Diego: Elsevier. Thybo, A.K., and M. Martens. 1999. Instrumental and sensory characterization of cooked potato texture. Journal of Texture Studies 30: 259278. Thybo, A.K., J.P. Mlgaard, and U. Kidmose. 2002. Effect of different organic growing conditions on quality of cooked potatoes. Journal of the Science of Food and Agriculture 82: 1218. Thybo, A.K., J. Christianson, K. Kaack, and M.A. Peterson. 2006. Effect of cultivars, wound healing, and storage on sensory quality and chemical components in pre-peeled potatoes. LWTFood Science and Technology 39: 166176. Troncoso, E., F. Pedreschi, and R.N. Ziga. 2009. Comparative study of physical and sensory properties of pre-treated potato slices

217 during vacuum and atmospheric frying. LWTFood Science and Technology 42: 187195. True, R.H., and T.M. Work. 1981. Sensory quality of Ontario potatoes compared with principal varieties grown in Maine. American Potato Journal 58: 375379. Ugawa, T., and K. Kurihara. 1994. Enhancement of canine taste responses to umami substances by salts. American Journal of Physiology 266: 944949. Ulrich, D., E. Hoberg, W. Neugebauer, H. Tiemann, and U. Darsow. 2000. Investigation of the boiled potato flavor by human sensory and instrumental methods. American Journal of Potato Research 77: 111117. Uppal, D.S. 1987. Varietal and environmental effect on the glycoalkaloid content of potato (Solanum tuberosum L.). Qualitas Plantarum 37: 333340. Vainionpaa, J., R. Kervinen, M. De Prado, E. Laurila, M. Kari, L. Mustonen, and R. Ahvaneainen. 2000. Exploration of storage and process tolerance of different potato cultivars using principal component and canonical correlation analyses. Journal of Food Engineering 44: 4761. Valkonen, J.P.T., M. Keskitalo, T. Vasara, and L. Pietila. 1996. Potato glycoalkaloids: A burden or a blessing? Critical Reviews in Plant Sciences 15: 120. van Dijk, C., M. Fischer, J. Holm, J.G. Beekhuizen, T. Stolle-Smits, and C. Boeriu. 2002. Texture of cooked potatoes (Solanum tuberosum). 1. Relationships between dry matter content, sensory-perceived texture, and near-infrared spectroscopy. Journal of Agricultural and Food Chemistry 50: 50825088. van Marle, J.T., R. van Der Vuurst De Vries, E.C. WIlkinson, and D. Yuksel. 1997a. Sensory evaluation of the texture of steam-cooked table potatoes. Potato Research 40: 7990. van Marle, J.T., T. Stolle-Smits, J. Donkers, C. van Dijk, A.G.J. Voragen, and K. Recourt. 1997b. Chemical and microscopic characterization of potato (Solanum tuberosum L.) cell walls during cooking. Journal of Agricultural and Food Chemistry 45: 5058. Wang, Y., and S.J. Kays. 2000. Contribution of volatile compounds to the characteristic aroma of baked Jewel sweetpotatoes. Journal of the American Society for Horticultural Science 125: 638643. Wang, Y., and S.J. Kays. 2003. Analytically directed flavor selection in breeding food crops. Journal of the American Society for Horticultural Science 128: 711720. Weiss, T.J. 1983. Food oils and their uses, 2nd ed. Westport: AVI Publishing Co., Inc. Winfield, M., D. Lloyd, W. Griffiths, J. Bradshaw, D. Muir, I. Nevison, and G. Bryan. 2005. Assessing organoleptic attributes of Solanum tuberosum and S. phureja potatoes. Aspects of Applied Biology 76: 127135. Woese, K., D. Lange, C. Boess, and K.W. Bgl. 1997. A comparison of organically and conventionally grown foodsResults of a review of the relevant literature. Journal of the Science of Food and Agriculture 74: 281293. Wszelaki, A.L., J.F. Delwiche, S.D. Walker, R.E. Liggett, J.C. Scheerens, and M.D. Kleinhenz. 2005. Sensory quality and mineral and glycoalkaloid concentrations in organically and conventionally grown redskin potatoes (Solanum tuberosum). Journal of the Science of Food and Agriculture 85: 720726.